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The Dryopteris Dilatata Complex in Macaronesia and the Iberian Peninsula

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The Dryopteris Dilatata Complex in Macaronesia and the Iberian Peninsula Acta Botánica Malacitana, 8: 59-72 Málaga, 1983 THE DRYOPTERIS DILATATA COMPLEX IN MACARONESIA AND THE IBERIAN PENINSULA M. GIBBY SUMMARY: Diploid and tetraploid species of the Dryopteris dilatata com- plex are found in Spain and Portugal. In an attempt to determine the re- lationships between species the cytology of both wild and synthetised hybrids has been investigated, and the results obtained so far will be described. RESUMEN: Se han encontrado especies diploides y tetraploides del complejo Dryopteris dilatata en España y Portugal. Para determinar el parentesco entre las especies se ha estudiado la citología de los híbridos tanto naturales como artificiales, exponiéndose finalmente los resultados obte- nidos. Ten species are at present recognised as belonging to the Dryop- teris dilatata (Hoffm.) A. Gray complex, eight of which can be found within the region of Macaronesia and the Iberian Peninsula. The other two species of the complex are native to eastern North America. Of the ten species of the complex, five are diploid, with 82 chromo- somes (2n = 82; see Plate 1) and five are tetraploid, with 2n = 164 (Plate 2). D. dilata itself is a tetraploid with an ovate-deltoid lamina that is mid to dark green in colour, and somewhat glandular, with spar- se stalked glands. The scales at the stipe base are triangular to linear in shape, usualy with a distinct broad dark central stripe. The rhizome is upright and the fronds have a lax growth form. An importan distinguishing character is the length of the lowest inner- most basiscopic pinnule which is usually equal in length to its neighbour; this distinguishes D. dilatata from some of the diploid species of the complex. D. dilatata has relatively large spinulose spores, 35-48 pm in length. The distribution of the species in Macaronesia and the Iberian Peninsula is described in Table 1. Only one species, the diploid D. aemula (Aiton) O. Kuntze, is found throughout the region. It is re- cognised by its triangular frond, dark brown or purplish stipe, dark concolorous scales, spineless spores, 31 -39 pm in length, up- curling pinnule margins and the presence of many pale green or white sessile glands on the lamina, which give the fern its charac- teristic smell of coumarin when dried. This is a common fern in the Azores and Madeira, but it has a much more restricted distribution 59 M. GIBBY in the Canary Islands, being recorded so far only from the El Cedro region of Gomera. In the Iberian Peninsula it is most frequent in Galicia. Several of the other species appear to be island endemics. D. azorica (Christ)) Alston (2n = 82; Plante 1) and D. crispifolia Rasbach, Reichstein & Vida (2n = 164; Plate 2) occur only on the Azores; D. azorica has been recorded from all the islands except Corvo and Graciosa (Gibby et al., 1977) whilst D. crispifolia is pro- bably limited to Pico and Faial. D. azorica has an ovate-deltate frond and the ultimate segments are broad and often touching which gives the frond a dense silhouette; glands are absent but the costae of the pinnules bear small brown scales, and the lowest innermost basiscopic pinnule is usually shorter than its neighbours, and often overlaps the rachis. The scales at the base of the stipe have a dark line, and the spore size is small, 28-38 pm in length. D. crispifolia is characterised by its very crisped appearance, which is quite dis- tinct from any other European Dryopteris. Scales on the stipe base are dark brown, sometimes showing a dark line, and long and nar- row in shape with a pointed tip. The lamina is covered with gree- nish white sessile glands, like D. aemula and the frond smells of coumarin. The midribs of the pinnae and pinnules bear small brown scales. Pinnules and pinnule segments are broad and often overlap- ping. The spore size is large, 35-45 pm . D. maderensis Alston (2n = 82) is a laurel forest species occurring only in Madeira. It is si- milar to D. azorica in having a small spore size (30-36 pm ), and a short lowest innermost basiscopic pinnule. The scales on the stipe base are dark brown and the lamina lacks glands but also lacks the small brown scales that are seen on the pinnules of D. azorica. The pinnules and pinnule segments of D. maderensis are narrower and not overlapping or touching, and both are deflexed from the mi- dribs; the ultimate teeth may upturn. Dryopteris guanchica Gibby & Jermy (2n = 164) has an ovate-del- tate lamina, distinctly stalked pinnules and lanceolate brown conco- lorous scales at the stipe base. On the lowest pairs of pinnae, the upper innermost pinnule is often distinctly smaller than its neigh- bours. The frond is usually flat, with upturned teeth, and the spo- res are large, 35-47 pm long. This species was first believed to be endemic to the Canary islands. It is recorded from Tenerife, Gomera and La Palma, although it is probably extinct in La Palma now. Mo- re recently it has been found to be present in the Iberian Peninsu- la. Amongst herbarium specimens from the Botanical Institute in Coimbra, Portugal, some specimens of D. guanchica were identified; these had been collected in the serra da Cintra, near Lisboa in 1839, probably by Welwitsch. During a visit to the same area in 1976 Dy- ce, Gibby and Jermy confirmed that D. guanchica is still growing at this site. Subsequently material of D. guanchica has been identi- fied from northern Portugal and north-west Spain. Dryopteris expansa (C. Presl) Fraser-Jenkins and Jermy (2n = 82) and D. carthusiana (Vill. ) H.P. Fuchs (2n = 164) have not been recorder from Macaronesia, and their distribution in the Iberian Pe- ninsula is limited mainly to the north-east and north-west. D expan- sa is similar to D. dilatata in morph'ology, but the segments are mo- re finely divided, the lamina is lighter green in colour and the sca- les are not usually so dark, being sometimes ginger and concolorous, or with a darker central stripe. The lowest innermost basiscopic pin- nule is usually long, often twice as long as its neighbour, in com- parison with D. dilatata where these are usually similar in length. As a result the lowest pinnae of D. expansa are distinctly triangu- lar in shape. The spores are large in D. expansa (35-48 pm ), slrni- Dryopteris dilatata complex lar in size to those of D. dilatata. D. carthusiana is characterised by pale brown concolorous stipe scales, and a creeping rhizome (un- like the other species of the complex, where the rhizome is upright); the fronds have an upright habit, a flat lamina without glands, and distinctive spinulose teeth on the pinnules. The stipe and lamina are usually equal in length, whereas in D. dilatata the stipe is usually half to two-thirds the length of the lamina. The spores of D. cart- thusiana are large, 35-47 pm in length. Records for D. dilatata throughout the region have long been con- fused, largely because it is only recently that D. azorica, D. ex- pansa, D. maderensis, D. crispifolia and D. guanchica have been recognised as separate species. D. dilatata does have a fairly wide distribution in the Iberian Peninsula , but its presence in Macarone- sia has been in doubt (Gibby et al. , 1977), However, some collec- tions of Dryopteris from Pico by Spgren are referable neither to D. aemula, D. azorica or D. crispifolia, and the morphology of these specimens looks very similar to that of D. dilatata from Europe. Re- cently Fraser-Jenkins has revisited the site of this material in Pico and collected living specimens. Root squashes of one plant show it to be tetraploid, with 164 chromosomes, and this, together with mor- phological similarities, suggests that it is indeed D. dilatata. The inter-relationships of these eight species within the complex have been determined by cytogenetic studies, based on analysis of meiosis in both wild and synthesised hybrids involving these and other Dryopteris species. The methods that were used in the synthe- sis of hybrids have been described in detail by Lovis (1968); meio- tic preparations were made by the aceto-carmine squash technique of Manton (1950). For mitotic preparations actively growing root tips were pre-:reated in a saturated solution of alphabromonaphthalene, fixed in 3:1, absolute alcohol: glacial acetic acid and stored in re- frigerator at -20°C. chromosome DIPLOID AA bp. AUTOTETRAPLOID AA AA doubling DIPLOID AA hybridisalion chromosome so. B mm- AA BB doubling A LLOT ETRAPLOI D DIPLOID B B Figure 1.- Diagrammatic representation of the formation of an autotetraploid and an allotetraploid. A and B represent non-homologous genomes. 61 M. GIBBY Tetraploids can be classified into basic types, autotetraploid and allotetraploid. The former is derived from a diploid by chromosome doubling, whereas the latter is der ived from two different diploid species, by hybridisation and chromosome duplication ( see Fig. 1 ) . A diploid can be represented as AA where A represents a chromosome set or genome ; in Dryopteris each genome consists of 41 chromosomes. The two chromosome sets in the diploid can pair at meiosis, forming 41 bivalents and so they are said to be homologous, and are repre- sented by the same letter. The autotetraploid has four homologous chromosome sets and is represented as AAAA (Fig. 1). The allotetra- ploid is designated AABB because it is derived from two different di- ploids, whose genomes are not homologous. In theory an autotetra- ploid should form quadrivalents during the first division of meiosis, and an allotretraploid should form only bivalents. However, autote- traploid ferns are normally bivalent - forming, probably as a result of some genetic control of chromosome pairing.
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